Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Heterobasidion annosum sensu lato
(Heterobasidion root rot)

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Datasheet

Heterobasidion annosum sensu lato (Heterobasidion root rot)

Summary

  • Last modified
  • 11 October 2017
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Natural Enemy
  • Preferred Scientific Name
  • Heterobasidion annosum sensu lato
  • Preferred Common Name
  • Heterobasidion root rot
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Fungi
  •     Phylum: Basidiomycota
  •       Subphylum: Agaricomycotina
  •         Class: Agaricomycetes

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Pictures

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PictureTitleCaptionCopyright
H. annosum causing death of young pine trees (Pinus spp.) in Thetford Forest, Norfolk, UK.
TitleSymptoms on young pine trees
CaptionH. annosum causing death of young pine trees (Pinus spp.) in Thetford Forest, Norfolk, UK.
CopyrightDavid S. Ingram
H. annosum causing death of young pine trees (Pinus spp.) in Thetford Forest, Norfolk, UK.
Symptoms on young pine treesH. annosum causing death of young pine trees (Pinus spp.) in Thetford Forest, Norfolk, UK.David S. Ingram

Identity

Top of page

Preferred Scientific Name

  • Heterobasidion annosum sensu lato (Fr.) Bref. 1888

Preferred Common Name

  • Heterobasidion root rot

Other Scientific Names

  • Fomes annosus (Fr.) Cooke 1885
  • Fomitopsis annosa (Fr.) P. Karst. 1881
  • Oedocephalum lineatum B.K. Bakshi 1950
  • Polyporus annosus Fr. 1821
  • Spiniger meineckellus (A.J. Olson) Stalpers 1974
  • Trametes radiciperda R. Hartig 1874
  • Ungulina annosa (Fr.) Pat. 1900

International Common Names

  • English: annosum root rot; annosus root rot; butt rot: conifers; conifers butt-rot; conifers heart rot; conifers red rot; Fomes root rot; heart rot: conifers; red rot: conifers; root rot: conifers; root rot: Hevea spp.
  • Spanish: podredumbre de los arboles resinosos
  • French: coeur rouge de l'epicea; le fomes; maladie du rond des pins; pourriture rouge des coniferes

Local Common Names

  • Germany: Heterobasidion Stamm- und Wurzelfäule; Heterobasidion-Fäule; Kernfäule; Rotfaeule: Fichte; Rotfäule: Fichte; Stockfaeule; Stockfäule; Wurzelfaeule; Wurzelfäule; Wurzelschwamm; Wurzelschwamm: Nadelhoelzer

EPPO code

  • HETEAN (Heterobasidion annosum)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Basidiomycota
  •             Subphylum: Agaricomycotina
  •                 Class: Agaricomycetes
  •                     Subclass: Agaricomycetidae
  •                         Order: Russulales
  •                             Family: Bondarzewiaceae
  •                                 Genus: Heterobasidion
  •                                     Species: Heterobasidion annosum sensu lato

Notes on Taxonomy and Nomenclature

Top of page Interfertility studies and DNA analyses have shown that the former Heterobasidion annosum sensu lato is a species complex consisting of at least four intersterile, or almost intersterile, taxa that show differences in distribution and host preference. As a consequence of these studies, Buchanan (1988) separated the non-pathogenic form of Heterobasidion occurring in Australia and adjacent regions from the Northern Hemisphere H. annosum as a new species, H. araucariae. Later, the European intersterility groups 'P', 'S' and 'F' of H. annosum sensu lato were recognized at species level by Niemelä and Korhonen (1998), with the names H. annosum sensu stricto, H. parviporum and H. abietinum, respectively.

These new species are treated separately in the Compendium, but the Northern Hemisphere taxa in the H. annosum complex are also treated as a collective species, H. annosum sensu lato, because their distribution is not well known. This complex includes the Eurasian species H. annosum sensu stricto, H. parviporum, H. abietinum and the North American intersterility groups 'P' and 'S'. The taxonomic relationships between the North American groups and the Eurasian species are not yet clear. In the Compendium, these groups are covered in the datasheets on H. annosum sensu stricto and H. parviporum, respectively.

All species and groups of the H. annosum complex show some sexual compatibility with other members of the complex in laboratory tests, but hybrids are very rare in nature (Gonthier et al., 2001). A case of hybridization has been reported in Europe (Schulze, 2000). Hybridization between the local S and P group appears to be more common in western North America. In greenhouse inoculation experiments, hybrid isolates showed lower pathogenicity than the parent groups (Garbelotto et al., 1998).

In addition to the Northern Hemisphere H. annosum sensu lato and Southern Hemisphere H. araucariae, the genus Heterobasidion includes another distinctly defined species, H. insulare, an almost non-pathogenic wood-decay fungus occurring in eastern and southern Asia. A number of other species have been described within this genus but their status as 'good' species has not been proved (Niemelä and Korhonen, 1998).

Description

Top of page Basidioma perennial, very irregular in shape, often imbricate and confluent, sesile with broad basal attachment, at times resupinate. Pileus 0-15 x 2-25 x 0·5-3 cm, convex, applanate, dimidiate or appressed-reflexed; upper surface grey-brown to bay, darkening with age, glabrescent, leaving a tuberculate-rugose crust, zonate; margin white, thin, acute. Context 0·2-1 cm thick, whitish, firm, corky to woody. Pore surface white to yellowish; pores irregular, circular to labyrinthoid, (1-) 2-3 per mm, 150-500 (-1000) µm diameter, dissepiments 40-120 µm thick, edge entire; tubes unevenly or distinctly stratified, 2-10 mm long in each layer, old tubes white stuffed. Basidiospores 3·5-5·5 x 3-4 (av. 4·5 x 3·5) µm, ovoid to broadly ellipsoid, hyaline, thin-walled, slightly asperulate, with a few guttulate contents. Basidia 9-13 x 5-7 µm, short clavate, 4-spored, without basal clamp. Cystidia absent. Hyphal system dimitic, non-agglutinated, with generative hyphae and skeletal hyphae. Generative hyphae 1·5-3 µm diameter, hyaline, thin-walled, freely branching, simple septate. Skeletal hyphae 1·5-5 µm diameter, hyaline, unbranched, of unlimited growth, thick-walled with narrow lumen, non-septate. Conidiophores produced in culture, oedocephaloid, vertical. Conidia 4·5-6·5 x 3-5·5 µm, non-septate, hyaline, ellipsoid to piriform, thick-walled (Pegler and Waterston, 1968).

Colonies growing rapidly on malt agar, 6-8 cm in 7 days, optimum temperature 26°C, strong reaction to tannic acid media (Campbell, 1938), mat floccose cottony, pale fawn, azonate, soon pulverulent due to conidial production (Cartwright and Findlay, 1946); cuticular cells and interlocking hyphae reported in some isolates forming pellicular crustose areas, test for extracellular oxidase negative (Nobles, 1965).

Pure cultures of H. annosum sensu lato originating from Europe or North America can be identified easily on the basis of typical conidiophores. Identification of cultures originating from eastern and southern Asia is more difficult because the local species H. insulare produces similar conidiophores, as does the Australian species H. araucariae.

Distribution

Top of page Some earlier records of H. annosum sensu lato from eastern and southern Asia may, in fact, represent Heterobasidion insulare, which can look very similar to H. annosum sensu lato, or other species of Heterobasidion that may occur in this region (Buchanan, 1988; Niemelä and Korhonen, 1998).

Distribution Table

Top of page

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

AfghanistanPresentNative Not invasive Buchanan, 1988
ChinaRestricted distributionEPPO, 2014
-JilinRestricted distributionNative Not invasive Dai et al., 2003
-SichuanPresentNative Not invasive CMI, 1980; EPPO, 2014
-TibetRestricted distributionNative Not invasive Mao, 1993
-XinjiangRestricted distributionNative Not invasive CMI, 1980; Dai et al., 2003
-YunnanPresentCMI, 1980; Dai et al., 2003; EPPO, 2014
IndiaRestricted distributionEPPO, 2014
-AssamPresentNative Not invasive CMI, 1980; EPPO, 2014
-Himachal PradeshPresentNative Not invasive CMI, 1980; EPPO, 2014
-Indian PunjabPresentNative Not invasive CMI, 1980; EPPO, 2014
-MeghalayaPresentNative Not invasive Roy and De, 1996
-Uttar PradeshPresentNative Not invasive CMI, 1980; EPPO, 2014
IsraelPresent Not invasive Binyamini, 1982
JapanWidespreadNative Not invasive CMI, 1980; Kobayashi et al., 1981; EPPO, 2014
KazakhstanRestricted distributionNative Not invasive Solov'ev, 1963
KyrgyzstanPresentNative Not invasive Dai et al., 2003; EPPO, 2014
MalaysiaRestricted distributionEPPO, 2014
-Peninsular MalaysiaPresentEPPO, 2014
MyanmarPresentNative Not invasive Hussain, 1952
NepalPresentNative Not invasive Adhikari, 1988
PakistanWidespreadHussain, 1952; CMI, 1980; EPPO, 2014
PhilippinesPresentNative Not invasive Eusebio, 1977
TurkeyWidespreadNative Not invasive CMI, 1980; EPPO, 2014
VietnamWidespreadCMI, 1980; Parmasto, 1986; EPPO, 2014

Africa

MoroccoWidespreadNative Not invasive CMI, 1980; EPPO, 2014

North America

CanadaRestricted distributionEPPO, 2014
-British ColumbiaPresentNative Not invasive CMI, 1980; EPPO, 2014
-Nova ScotiaPresentPunter, 1970; CMI, 1980; EPPO, 2014
-OntarioPresentNative Invasive Punter, 1970; CMI, 1980; EPPO, 2014
-QuebecRestricted distributionNative Invasive Laflamme and Blais, 1995
MexicoWidespreadNative Not invasive CMI, 1980; EPPO, 2014
USAWidespreadCMI, 1980; EPPO, 2014
-AlabamaWidespreadNative Not invasive Hodges, 1969
-AlaskaRestricted distributionNative Not invasive CMI, 1980
-ArizonaRestricted distributionNative Not invasive Gilbertson et al., 1974; USDA Forest Service, 2002
-ArkansasRestricted distributionNative Not invasive Wilson, 1963
-CaliforniaWidespreadNative Not invasive USDA Forest Service, 2002
-ColoradoWidespreadNative Not invasive James and Goheen, 1981; USDA Forest Service, 2002
-ConnecticutWidespreadNative Not invasive Stoddard et al., 1939
-DelawareRestricted distributionNative Not invasive Hartley, 1910; USDA Forest Service, 1997
-FloridaWidespreadNative Not invasive USDA Forest Service, 2002
-GeorgiaWidespreadNative Not invasive Froelich et al., 1977; USDA Forest Service, 2002
-IdahoWidespreadNative Not invasive Williams, 1989; USDA Forest Service, 2002
-IllinoisRestricted distributionNative Not invasive Hanson and Lautz, 1971
-IndianaRestricted distributionNative Not invasive Williams, 1959
-IowaRestricted distributionNative Not invasive Anderson et al., 1976
-KentuckyRestricted distributionNative Not invasive Kentucky Division of Forestry, 2003
-LouisianaRestricted distributionNative Not invasive Froelich et al., 1977
-MaineRestricted distributionNative Not invasive Shigo, 1975
-MarylandRestricted distributionNative Not invasive Little et al., 1968
-MassachusettsRestricted distributionNative Not invasive Sinclair, 1964
-MichiganRestricted distributionNative Not invasive Strong and Lemmien, 1964
-MinnesotaRestricted distributionNative Not invasive Gilbertson and Lombard, 1976
-MississippiRestricted distributionNative Not invasive Froelich et al., 1977
-MissouriWidespreadNative Not invasive Berry and Dooling, 1962
-MontanaWidespreadNative Not invasive Williams, 1989; USDA Forest Service, 2002
-NebraskaRestricted distributionNative Not invasive USDA Forest Service, 2002
-NevadaWidespreadNative Not invasive Tegethoff, 1973; USDA Forest Service, 2002
-New HampshireWidespreadNative Not invasive Miller, 1960
-New JerseyPresentNative Not invasive Sinclair, 1964
-New MexicoRestricted distributionNative Not invasive Gilbertson et al., 1975; USDA Forest Service, 2002
-New YorkWidespreadNative Not invasive Sinclair, 1964
-North CarolinaWidespreadNative Not invasive Jacobi et al., 1980; USDA Forest Service, 2000
-OhioRestricted distributionNative Not invasive Berry, 1968
-OregonWidespreadNative Not invasive USDA Forest Service, 2002
-PennsylvaniaRestricted distributionNative Not invasive Stambaugh et al., 1962
-Rhode IslandWidespreadNative Not invasive Hadfield, 1970
-South CarolinaWidespreadNative Not invasive Powers and Boyce, 1961; USDA Forest Service, 2002
-South DakotaPresentNative Not invasive Sinclair, 1964
-TennesseeWidespreadNative Not invasive Applegate, 1971
-TexasRestricted distributionNative Not invasive Mason, 1969; USDA Forest Service, 2000
-UtahRestricted distributionNative Not invasive Williams, 1989; USDA Forest Service, 2002
-VermontWidespreadNative Not invasive Chase and Ullrich, 1983
-VirginiaWidespreadNative Not invasive Webb et al., 1981
-WashingtonWidespreadNative Not invasive USDA Forest Service, 2002
-West VirginiaWidespreadNative Not invasive Ginns and Gillespie, 1962
-WisconsinRestricted distributionNative Not invasive Stanosz and Guthmiller, 1995; USDA Forest Service, 2002
-WyomingRestricted distributionNative Not invasive Tegethoff, 1973; USDA Forest Service, 2002

Central America and Caribbean

CubaWidespreadNative Not invasive CMI, 1980; EPPO, 2014
Dominican RepublicWidespreadNative Not invasive CMI, 1980; EPPO, 2014
GuatemalaPresentNative Not invasive Lowe, 1957
HondurasRestricted distributionNative Not invasive
JamaicaWidespreadNative Not invasive CMI, 1980; Greig and Foster, 1982; EPPO, 2014

Europe

AlbaniaRestricted distributionNative Not invasive Lushaj, 2001
AustriaWidespreadNative Not invasive CMI, 1980; EPPO, 2014
BelarusPresentEPPO, 2014
BelgiumWidespreadNative Not invasive CMI, 1980; EPPO, 2014
Bosnia-HercegovinaPresentNative Not invasive Marinkovic, 1980
BulgariaRestricted distributionNative1907 Not invasive CMI, 1980; Rosnev and Petkov, 1986; EPPO, 2014
CroatiaWidespreadNative Not invasive Halambek et al., 1996
Czech RepublicWidespreadNative Not invasive CMI, 1980; Kotlaba, 1984; EPPO, 2014
Czechoslovakia (former)Widespread****CMI, 1980; EPPO, 2014
DenmarkWidespreadNative Not invasive CMI, 1980; Thomsen, 1994; EPPO, 2014
EstoniaPresentNative Not invasive Hanso and Hanso, 2003; EPPO, 2014
Faroe IslandsPresentVesterholt and Pedersen, 1995
FinlandWidespreadNative Not invasive CMI, 1980; EPPO, 2014
FranceWidespreadNative Not invasive CMI, 1980; EPPO, 2014
GermanyWidespreadNative**** Not invasive CMI, 1980; EPPO, 2014
GreeceEradicatedNative Not invasive Tsopelas and Korhonen, 1996
HungaryWidespreadNative Not invasive CMI, 1980; EPPO, 2014
IrelandWidespreadNative Not invasive CMI, 1980; EPPO, 2014
ItalyWidespreadNative Not invasive CMI, 1980; EPPO, 2014
LatviaPresentNative Not invasive Grantina et al., 2000; EPPO, 2014
LithuaniaPresentNative Not invasive Vasiliauskas et al., 2002; EPPO, 2014
MacedoniaPresentNative Not invasive Marinkovic, 1980
MoldovaRestricted distributionNative Not invasive Gorova, 1980
NetherlandsWidespreadNative Not invasive CMI, 1980; EPPO, 2014
NorwayWidespreadNative**** Not invasive CMI, 1980; EPPO, 2014
PolandWidespreadNative Not invasive CMI, 1980; EPPO, 2014
PortugalWidespreadNative Not invasive CMI, 1980; EPPO, 2014
-MadeiraPresentKorhonen, 1978
RomaniaWidespreadNative Not invasive CMI, 1980; EPPO, 2014
Russian FederationRestricted distributionEPPO, 2014
-Eastern SiberiaPresentEPPO, 2014
-Russia (Europe)PresentNative Not invasive EPPO, 2014
-Russian Far EastPresentNative Not invasive CMI, 1980; Morozova and Tkacz, 1997; EPPO, 2014
-SiberiaPresentNative Not invasive CMI, 1980; Negrutskii, 1986
-Western SiberiaPresentEPPO, 2014
SerbiaWidespreadEPPO, 2014
SlovakiaWidespreadNative Not invasive Kotlaba, 1984
SloveniaWidespreadNative Not invasive Munda et al., 1998
SpainWidespreadNative Not invasive CMI, 1980; EPPO, 2014
SwedenWidespreadNative**** Not invasive CMI, 1980; EPPO, 2014
SwitzerlandWidespreadNative**** Not invasive CMI, 1980; EPPO, 2014
UKRestricted distributionNative Not invasive EPPO, 2014
-England and WalesRestricted distributionNative Not invasive EPPO, 2014
-Northern IrelandRestricted distributionNative Not invasive EPPO, 2014
-ScotlandRestricted distributionNative Not invasive EPPO, 2014
UkrainePresentNative Not invasive EPPO, 2014
Yugoslavia (Serbia and Montenegro)WidespreadNative Not invasive CMI, 1980; Tomanic et al., 2000

Oceania

AustraliaRestricted distributionEPPO, 2014
-New South WalesPresentEPPO, 2014
-QueenslandPresentEPPO, 2014
FijiWidespreadEPPO, 2014
New ZealandWidespreadEPPO, 2014
Papua New GuineaWidespreadEPPO, 2014

History of Introduction and Spread

Top of page There are no proven cases of introduction of H. annosum sensu lato.

Hosts/Species Affected

Top of page The host range of H. annosum sensu lato includes a wide range of Northern Hemisphere conifers, less commonly broadleaved trees and other woody plants, and rarely non-woody plants.

For further information on hosts, see Sinclair (1964), Greig (1976), Delatour (1977), Wagn (1980) and Webb and Alexander (1985).

Growth Stages

Top of page Vegetative growing stage

Symptoms

Top of page H. annosum sensu lato causes root decay in all hosts, but the extent of stem decay varies according to the host species (Greig, 1998). In resinous conifers, like most pine species, infection causes profuse resin excretion on the roots and at the base of the stem. This excretion blocks the flow of fluids in the stem. The tree dies relatively soon afterwards; the green crown turns brown and dies simultaneously.

In coniferous genera such as Picea, Abies, Larix, Pseudotsuga and Tsuga, decay rises up the stem, usually as a heart rot. The tree may be asymptomatic for decades after infection. External symptoms only appear at an advanced stage of decay and include reduced growth, defoliation of the crown, and resin exudations on the stem. Wind-thrown trees with decayed roots or open gaps in the stand often indicate the presence of Heterobasidion root rot in the forest. In the field, Heterobasidion root rot can only be identified with assurance by the presence of basidiocarps, which usually hide under decayed roots, moss cover at the stem base of dead trees, in hollow stumps, or under the fallen trunks of diseased trees.

Stem decay caused by Heterobasidion remains relatively hard for long time and is usually light brown in colour. The first stage of decay often has a violet coloration. Advanced decay contains small, white cellulose pockets ('white pocket rot'), often with black spots.

List of Symptoms/Signs

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SignLife StagesType
Roots / rot of wood
Stems / discoloration of bark

Biology and Ecology

Top of page Members of the H. annosum complex are root rot fungi that spread vegetatively via root contact and grafts, forming distinct disease centres in the tree stand (Stenlid and Redfern, 1998). These disease centres begin with the infection of freshly cut conifer stumps or wounds on the roots or (more seldom) stems of living conifers by spores of the fungus (Redfern and Stenlid, 1998). In large stumps the fungus can remain alive and infectious for decades after felling and thus forms a threat for the next conifer generation (Piri, 1996).

Heterobasidion root rot is most damaging on soils with a high pH and in afforestations on former agricultural soil. The frequency of rot is very dependent on forest management practices.

Transmission

Heterobasidion spores may become airborne when daily temperatures rise above 0°C, and their number can be substantial above 10°C (Redfern and Stenlid, 1998). Long, dry periods will interrupt spore production by the fungus, and high temperatures on stump surfaces will prevent infection (Driver and Ginns, 1969). Spores deposited on the forest soil are carried down by rainwater and may remain infective for several months (Kuhlman, 1966). Spores may be introduced into disease-free areas on nursery seedlings (Jorgensen, 1961).

Physiological Specialization

Different members of the H. annosum complex show distinct physiological differentiation: for example, in enzymatic properties (Maijala et al., 2003) and wood-decaying ability (Daniel et al., 1998).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Armillaria mellea Pathogen
Bjerkandera adusta Pathogen
Bjerkandera adusta Pathogen
Fomitopsis pinicola Pathogen
Gloeophyllum saepiarium Pathogen
Hirschioporus abietinus Pathogen
Hypholoma capnoides Pathogen
Hypocrea rufa Mycoparasite Italy
Macrolepiota procera Pathogen
Macrolepiota rachodes Pathogen
Neonectria fuckeliana Pathogen
Oidiodendron maius Pathogen
Phlebia gigantea Pathogen
Pleurotus ostreatus Antagonist
Polyporus benzoinus Pathogen
Resinicium bicolor Pathogen
Scytalidium album Pathogen
Scytalidium lignicola Pathogen
Sistotrema brinckmannii Pathogen
Tolypocladium niveum Pathogen

Means of Movement and Dispersal

Top of page Natural Dispersal

The great majority of spores fall within a few hundred metres of the source (Möykkynen et al., 1997), but viable, airborne spores have been found 300 km from the nearest source (Rishbeth, 1959). Conidia may also have some role in the aerial dispersal of Heterobasidion spores (Hsiang et al., 1989; Möykkynen, 1997).

Vector Transmission

Insects and other animals can carry spores of Heterobasidion but appear to be of minor importance in spreading the disease (Otrosina and Cobb, 1989).

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Roots hyphae Yes Yes Pest or symptoms usually invisible
Stems (above ground)/Shoots/Trunks/Branches fruiting bodies; hyphae Yes Yes Pest or symptoms usually visible to the naked eye
Wood
Plant parts not known to carry the pest in trade/transport
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Leaves
Seedlings/Micropropagated plants
True seeds (inc. grain)

Wood Packaging

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Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood
Solid wood packing material with bark
Solid wood packing material without bark

Prevention and Control

Top of page In healthy and slightly diseased stands in risky areas it is important to prevent infection by spores. This is done by performing cuttings during the seasons (cold winter or hot summer) when the risk of spore infection is low or absent. For cuttings carried out during the more risky warm season, the cut surface of fresh stumps can be treated with a protectant to prevent infection. The most commonly used protectants are urea and borax (Pratt et al., 1998) and the competitive fungus Phlebia gigantea (Holdenrieder and Greig, 1998). Care should be taken to avoid wounding the roots during logging operations, especially when cuttings are carried out during the warm season. Wide spacing and minimizing the number of thinnings during rotation will reduce the risk of infection.

Little can be done to control the fungus in diseased stands. The only real possibility is to shorten the rotation time in heavily diseased stands.

The extraction of infected stumps before regeneration essentially reduces the risk of infection in the next generation of susceptible conifers. The spread of the disease from stumps can also be reduced by favouring the regeneration of resistant tree species close to decayed stumps. Heterobasidion infection can be cleared from a site by cultivating a rotation of a resistant tree species. Current knowledge suggests that pure broadleaved tree stands are practically resistant. Susceptible conifers should be avoided or cultivated in mixture with resistant trees in afforestations on former agricultural soils (Korhonen et al., 1998b).

References

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Adhikari MK, 1988. Polypores (wood-rotting fungi) of Nepal. Banko Janakari Kathmandu, Nepal; Forest Research and Information Centre, Forest Survey and Research Office, Department of Forests, 2(1):9-20

Anderson RL; Ritter W; Witmer RC, 1976. Fomes annosus found on eastern white pine in Iowa. Plant Disease Reporter, 60:981-984.

Anon., 1959. Reports on forest research for the years ended March, 1957, March, 1958. London, UK: HMSO.

Anon., 1985. Taxonomy and ecology of wood-destroying fungi. Rep. For. Prod. Aust. 1965/66, 1966 (39).

Anon., 2004. Cylindroselloides dybasi, Hall in Nova Scotia. World Wide Web page at http://www.chebucto.ns.ca/Environment/NHR/cylindroselloides.html.

Applegate HW, 1971. Annosus root rot mortality in once-thinned loblolly pine plantations in Tennessee. Plant Disease Reporter, 55:625-627.

Bakshi BK, 1950. Fungi associated with ambrosia beetles in Great Britain. Transactions of the British Mycological Society, 33(1-2):111-120.

Bakshi BK; Sen M; Singh B, 1970. Cultural diagnosis of Indian Polyporaceae. II. Genera Fomes and Trametes. Indian Forest Records, 2:245-276.

Barnard EL; Gilly SP; Dixon WN, 1991. Incidence of Heterobasidion annosum and other root-infecting fungi in residual stumps and roots in thinned slash pine plantations in Florida. Plant Disease, 75(8):823-828

Berry FH, 1968. Spread of Fomes annosus root rot in thinned shortleaf pine plantations. USDA Forest Service, Res. Note NE-87, 4 pp.

Berry FH; Bretz TW, 1964. Urea and other chemicals effective against colonization of Shortleaf Pine stumps by Fomes annosus in Missouri. Plant Disease Reporter, 48(11):886-887.

Berry FH; Dooling OJ, 1962. Fomes annosus on shortleaf pine in Missouri. Plant Disease Reporter, 46:521-522.

Binyamini N, 1982. Host index for Israeli wood-rotting fungi. Tel Aviv University, Tel Aviv, 30 pp.

Buchanan PK, 1988. A new species of Heterobasidion (Polyporaceae) from Australasia. Mycotaxon, 32:325-337

Campbell WA, 1938. The cultural characteristics of the species of Fomes. Bulletin Torrey botanical Club, 65(1):31-69.

Capretti P; Barzanti GP; Cech T; Tomiczek C, 1998. Intersterility groups (ISG) of Heterobasidion annosum in the Italian Alpine region and Austria. In: Delatour C, Guillaumin J-J, Lung-Escarmant B, Marçais B, eds. Root and Butt Rots of Forest Trees. 9th International Conference on Root and Butt Rots. Carcans-Maubuisson, France. INRA Editions, Les Colloques, no. 89: 437. (Abstr.).

Capretti P; Barzanti GP; Luisi N; Puddu A, 1998. Group dying of Silver fir (Abies alba) by Heterobasidion annosum in Central and Southern Italy. In: Delatour C, Guillaumin J-J, Lung-Escarmant B, Marçais B, eds. Root and Butt Rots of Forest Trees. 9th International Conference on Root and Butt Rots. Carcans-Maubuisson, France. INRA Editions, Les Colloques, no. 89: 440. (Abstr.).

Capretti P; Goggioli V; Mugnai L, 1994. Intersterility groups of Heterobasidion annosum in Italy: Distribution, hosts and pathogenicity tests. In: Johansson M, Stenlid J, ed. Proceedings of the Eighth International Conference on Root and Butt Rots. Wik, Sweden and Haikko, Finland. Swedish Univ. of Agric. Sci., Uppsala, Sweden, 218-226.

Capretti P; Korhonen K; Mugnai L; Romagnoli C, 1990. An intersterility group of Heterobasidion annosum specialized to Abies alba. European Journal of Forest Pathology, 20(4):231-240

Capretti P; Tegli S; Lakomy P; Zamponi L, 2003. Genetic variation in Heterobasidion abietinum (H. annosum F group) population. In: Laflamme G, Bérubé JA, Bussières G, eds. Root and butt rots of forest trees. 10th International Conference on Root and Butt Rots, Québec City, Canada. Canadian Forest Service, Laurentian Forestry Centre, Information Report LAU-X-126:293-295.

Cartwright K St G; Findlay WPK, 1946. Decay of Timber and its Prevention. London, UK: HMSO.

Certini G; Corti G; Ugolini FC, 2000. Influence of soil properties on the mortality of silver fir in Tuscany, Italy. Forstwissenschaftliches Centralblatt, 119(6):323-331; 25 ref.

Chang TT; Hsieh HJ; Chang RJ; Fu CS, 1999. Common tree diseases in Taiwan. Taiwan Forestry Research Institute. (In Chinese).

Chase TE; Ullrich RC, 1983. Sexuality, distribution, and dispersal of Heterobasidion annosum in pine plantations of Vermont. Mycologia, 75(5):825-831

Chase TE; Ullrich RC, 1990. Genetic basis of biological species in Heterobasidion annosum: Mendelian determinants. Mycologia, 82(1):67-72

Chase TE; Ullrich RC; Korhonen K, 1985. Homothallic isolates of Heterobasidion annosum. Mycologia, 77(6):975-977

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